_ -, ' ' S / STATUS OF THE GENUS (: EULIMIDAE), */eV WITH A DESCRIPTION OF A NEW SPECIES ‘

William G. Lyons

Abstract.—Monotypie genera Spiroclimax Mörch, 1875, andAthleenia Bartsch, 1946, are later synonyms of Oceanida DeFolin, 1871; type-species of both genera are identical to O. graduata, the type-species ofOceanida. All were described from the tropical western Atlantic. Oceanclia inglei n. sp. from Florida and North Carolina differs from O. graduata by lacking tabulate shoulders at posterior angles of spiral whorls. Differences be­ tween Eulimidae (preferred to Melanellidae) and Stiliferidae are unclear, indicating the latter should be combined with the former family.

Three originally monotypic genera have been erected to contain the single species most recently known as Athleenia burryi Bartsch, 1946. The earliest available name for the genus appears to be Oceanida DeFolin, 1871. Another species, presently known from Florida and North Carolina, is previously unnamed and is described herein. Specimens were examined from collections of the Plourglass Cruises off west Florida, in samples taken during faunal studies near the Florida Power and Light nuclear facility at Hutchinson Island, Florida east coast, and in other programs conducted by the Florida Department of Natural Re­ sources (FDNR) Marine Research Laboratory. Most specimens are housed in the FDNR invertebrate reference collection (catalogue prefix FSBC I) at St. Petersburg. Other specimens were examined from collections of the National Museum of Natural History (USNM), Washington, D.C., the Steger collection, now in the Delaware Museum of Natural History (DMNH), Greenville, Delaware, and additional material collected at Hutchinson Island by Applied Biology, Inc., Jensen Beach, Florida. Speci­ mens were deposited in molluscan collections at the British Museum (Nat­ ural History) (BMNH), London, and the Museum of Comparative Zoology (MCZ), Harvard University, Cambridge, Massachusetts.

Genus Oceanida DeFolin, 1871 Oceanida DeFolin, 1871:264 (type-speciesOceanida graduata DeFolin, 1871, by monotypy).—Kisch, 1959:107. Spiroclimax Mörch, 1875:168 (type-speciesSpiroclimax scalaris Mörch, 1875, by monotypy). Chrysallida (Oceanida): Thiele, 1929:232. Odostomia (Spiroclimax): Thiele, 1929:235. 540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Athleenia Bartsch, 1946:30 (type-speciesAthleenia burryi Bartsch, 1946, by monotypy).—Abbott, 1974:130.

Diagnosis.—Shells small, with maximum lengths slightly less than 3.0 mm, glassy, smooth. Embryonic whorls tapered, rather slender, of about 3H-4 whorls, each bearing many very fine axial riblets. Postembryonic whorls broader than long, expanding rather rapidly, constricted at sutures, with about one weak growth interruption per whorl. Apertures tear-shaped, bordered by simple columella and outer lip. Remarks.— Oceanida was introduced without formal description to con­ tain O. graduata, a species described from Guadeloupe, West Indies, in the same paper by DeFolin (1871). No higher classification was suggested by DeFolin, but Thiele (1929) considered Oceanida a subgenus of the pyrami- dellid genus Chrysallida Carpenter, 1857. Kisch (1959) treatedOceanida as a genus of Chemnitzidae (= Pyramidellidae) in a review of DeFolin’s taxa of that family. Both Thiele and Kisch recognized O. graduata as the type- species of Oceanida. Four years after DeFolin introduced Oceanida, Mörch (1875) erected Spiroclimax for S. scalaris, described without illustration from a single specimen collected at St. Thomas, West Indies. Mörch considered Spiro­ climax a genus of Pyramidellidae. Thiele (1929:235, fig. 240) subsequently illustrated S. scalaris as the type-species ofSpiroclimax, which he con­ sidered a subgenus of Odostomia Fleming, 1817, another pyramidellid genus. Like Oceanida, Spiroclimax seems to have escaped further notice as a component of western Atlantic fauna. More recently, Bartsch (1946) proposedAthleenia to contain A. burryi, which he thought to be a new species from deep water off south Florida. He assigned the genus to Stiliferidae. At my request, Dr. Anders Waren examined the types of Oceanida graduata and Spiroclimax scalaris at the Muséum National d’Histoire Nat­ urelle, Paris, and the Universitetets Zoologiske Museum, Copenhagen, re­ spectively. We have both examined the holotype Athleenia of burryi (USNM 573616). According to Dr. Waren (personal communication), type-specimens of all three proposed names are identical and constitute a single species. Assignment of Oceanida to Pyramidellidae by Mörch (as Spiroclimax), Thiele, and Kisch seems inappropriate as the shells bear no features relating them to that family. Abbott (1974) followed Bartsch in placing Athleenia in Stiliferidae, but also suggested that Rissoidae might be more appropriate, apparently noting the same“forma Rissoae vitreae” mentioned by Mörch in his description of Spiroclimax. I have found no genera of Rissoidae re­ sembling Oceanida, but aperture and spire shape are suggestive of certain Rissoinidae, especially Crepitacella Guppy, 1867. The resemblance is VOLUME 91, NUMBER 2 541 superficial, however; species ofCrepitacella bear marked axial sculpture, a broad anal fasciole, and a strong basal fold extending into the aperture, all lacking in Oceanida. Shells of Oceanida seem similar to those of Turveria Berry, 1956, a monotypic genus erected in Stiliferidae to contain T. encopendema, a species parasitic on Encope echinoids in western Mexico. The species was il­ lustrated in two line drawings by Berry (1956: fig. 2) and another, probably redrawn from Berry, in Keen (1971), but I have been unable to obtain shells or photographs for comparison. Mörch, Thiele, Berry, and Abbott, in aforementioned papers, assigned synonyms of Oceanida and species in genera similar to that genus to Stiliferidae. Bartsch (1917) included Broderip and Sowerby, 1832, and other customarily stiliferid genera in his monograph of east Pacific Melanellidae, but later assigned Athleenia to Stiliferidae. Keen (1971) continued to include east Pacific stiliferid genera in Melanellidae. Most members of both families are parasitic on . Typical genera (e.g., Stilifer, ) are obviously disparate, the first having thin, rotund shells with mucronate embryonic whorls, while the other has slender, elongate, relatively thick shells with glassy, polished surfaces and evenly tapered embryonic whorls. Genera differing more than slightly from the Melanella design (e.g., Kohler and Vaney, 1908; A. Adams, 1860; “Athleenia” Bartsch, 1946) tend to be assigned to Stiliferidae. Berry (1956) seemed equally perplexed regarding familial assignment for his Turveria, opting for Stiliferidae because the shell pos­ sessed a “papilliform apex.” The apex he figured, however, resembles those found on Oceanida species, which seem little more related to Stilifer than to some melanellids. Problems have also arisen in selecting the proper name for the family containing Melanella Bowdich, 1822, primarily because of confusion re­ garding the supposed synonymous relationship between that genus and Risso, 1826, resulting in replacement of Eulimidae H. and A. Adams, 1853 (fide Golikov and Starobogatov, 1975) with Melanellidae Bartsch, 1917 (fide Abbott, 1974). Little would be accomplished by enumerating the many references maintaining this synonymy, but careful attention to dif­ ferences separating genera in this family indicates that Melanella and Eulima are not synonymous (see Lyons, 1977, for examples of both genera). However, given that such synonymy could be demonstrated, Eulimidae remains the correct name for the family even though its type- genus was proposed later than that of Melanellidae (see International Code of Zoological Nomenclature, XV International Congress, 1964, Art. 64). Mayr (1969:356) argues effectively for retention of early familial appellations regardless of subsequent type-genus changes. Because no well defined boundaries seem to exist between Eulimidae and 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Stiliferidae, it would appear that they should be combined. Since Stiliferidae (originally Styliferidae) H. and A. Adams, 1853 fide ( Abbott, 1974) and Eulimidae were described at the same time, subsequent selection is a matter of reviewer preference. The more diverse and abundant Eulimidae is herein preferred. Oceanida graduata DeFolin, 1871 Figs. 1-3 Oceanida graduata DeFolin, 1871:264, pi. 24, fig. 6.—Kisch, 1959:107. Spiroclimax scalaris Mörch, 1875:168. Chrysallida (Oceanida) graduata: Thiele, 1929:232. Odostomia (Spiroclimax) scalaris: Thiele, 1929:235, fig. 240. Athleenia burryi Bartsch, 1946:30, fig. 1.—Abbott, 1974:131, fig. 1433. Description.—Shell small, to about 2.8 nun length, with approximately 3% very finely ribbed embryonic whorls, followed by approximately 5% smooth, translucent, posteriorly tabulated postembryonic whorls. Aperture extended and rounded anteriorly; outer lip slightly compressed, extend­ ing nearly parallel to longitudinal axis of shell. Material examined.—FLORIDA: Holotype of Athleenia burryi, 2.1 mm; off Carysfort Reef Light, Monroe County, 400 ft (122 m); USNM 573616.— 1, 2.8 mm; % mi off Cape Florida Light, Dade County, 6 m; FSBC I 18166.—BAHAMAS: 1, 2.2 mm; Gibson Cay, Andros, beach drift; FSBC I 11748.—2, 2.1, 2.7 mm; South Bimini, beach drift; FSBC I 18164.— 1, 2.3 mm; Fernandez Bay, Cat Island, beach drift; FSBC I 18165. Remarks.—The holotype of Athleenia burryi is a dead shell described from a depth of 400 ft (122 m), but specimens I have seen from near Cape Florida and from beach drift at three Bahamian Islands indicate that the species probably dwells in shallows of the tropical western Atlantic. It is known to occur southward to St. Thomas, Virgin Islands (Mörch, 1875) and Guadeloupe (DeFolin, 1871).

Oceanida inglei, new species Fig. 4 Description.—Shell small, to about 2.7 mm, glassy, transluscent, fading to white when dead. Protoconch of about 4 glassy, transparent, yellow whorls marked by many very fine axial riblets or lines. Postembryonic whorls 4L- 5, expanding anteriorly, moderately rounded laterally, terminating abruptly posteriorly; anterior parts of spiral whorls seen through overlapping posterior portions of subsequent whorls as single, posterior spiral lines; each whorl marked by single axial growth line; penultimate whorl elongate, merging smoothly with base. Aperture tear-shaped, rounded, expanded anteriorly, constricted posteriorly; outer lip thin, simple, rounded; inner lip VOLUME 91, NUMBER 2 543

Figs. .1-4. Western Atlantic Oceanida. 1, Original illustration of Oceanida graduata, after DeFolin (1871: fig. 6); 2, Spiroclimax scalaris [ = O. graduata], after Thiele (1929: fig. 240); 3, O. graduata, 2.8 mm, from off Cape Florida, FSBC I 18166; 4,Oceanida inglei, n. sp., holotype, 2.7 mm, from off Hutchinson Island, Florida, USNM 77.1864. All 2 0 x - slightly elevated anteriorly, forming narrow pseudoumbilicus at inter­ section with base. H olotype.—Length 2.7 mm; off Hutchinson Island, St. Lucie County, Florida, 9.7 m depth; USNM 771864. Paratypes.—FLORIDA: 1, 1.9 mm; Florida Middle Ground in north­ eastern Gulf of Mexico, 28°38.1'N, 84°16.3'W, 28.6 m; FSBC I 18163.— 3, 2.4, 2.6 mm, fragment; Hourglass Station A, 27°35'N, 82°50'W, off Egmont Key, Pinellas County, 6 m; FSBC I 11747.—1, 2.6 mm; off Hutchin­ son Island, 11.3 m; FSBC I 11746.—1, 2.7 mm; off Hutchinson Island, 11.0 m; FSBC I 11745.— 1, 2.4 mm; off Plutchinson Island, 10.4 m; USNM 771865.—1, 2.7 mm; same data; MCZ 288503.—1, 2.6 mm; off Hutchinson Island, 10.7 m; BM NH 197813.— 1, 2.2 mm; off St. Augustine, St. Johns County, 38 m; DM NII 121798.—NORTH CAROLINA: 1, 2.1 mm; 12 mi east of Frying Pan Shoal, 22 m; USNM 82989. Etym ology.—The species is named for Robert M. Ingle, former Director of the Florida Department of Natural Resources Marine Research Lab­ oratory, who authorized and encouraged the Hutchinson Island faunal study. Remarks.—Oceanida inglei resembles 0. graduata in size, embryonic sculpture, shell surface texture, and general outline, but lacks the sharp, 544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON posterior shoulder angulations characteristic of the latter species. In addi­ tion, the aperture of O. inglei is slightly more constricted anteriorly than is that of its congener. The species are otherwise nearly identical. Most specimens of the new species were collected from coastal areas of east and west Florida, and were accompanied by fauna of warm temperate, rather than tropical, affinity. The only apparently tropical association yet noted for the species is at the Florida Middle Ground, where Caribbean species predominate (Lyons, 1976; Turgeon and Lyons, 1978).

Acknowledgments Dr. Anders Waren, University of Göteborg, Sweden, graciously examined types at two European museums; without his assistance, this study would not have been possible. Drs. Joseph Rosewater (USNM), R. Tucker Abbott (DMNH), and Donald R. Moore, Rosenstiel School of Marine and At­ mospheric Science, Miami, Florida, Mrs. Barbara Albert Steger, Tampa, Florida, and Robert M. Gallagher, Applied Biology, Inc., Jensen Beach. Florida, all loaned specimens for study. Dr. Moore also brought to my attention DeFolin s report. Mrs. Sally D. Kaicher, St. Petersburg, Florida, provided photographic assistance. All are gratefully thanked. The specimen from Andros, Bahamas, was collected during cruise G-7124 of the M/V Gerda, RSMAS research vessel.

Literature Cited

Abbott, R. T. 1974. American seashells, 2nd ed. 663 pp., 24 pis. Van Nostrand Reinhold Co., New York. Bartsch, P. 1917. A monograph of west American melanellid mollusks. Proc. U.S. Nat. Mus. 53(2207):295-356, 16 pis. ------. 1946. A new genus and species of minute marine stiliferid mollusk from Florida. J. Wash. Acad. Sei. 36(1): 30, 1 fig. Berry, S. S. 1956. A new west Mexican prosobranch mollusk parasitic on echinoids. Am. Midi. Nat. 56(2): 355-357. DeFolin, L. 1871. Chap. XXI. Nouveau supplément aux Mollusques de la Pointe-à- Pitre. Pp. 263-265, pi. 24in Les Fonds de la Mer. Golikov, A. N., and Y. I. Starobogatov. 1975. Systematics of prosobranch gastropods. Malacologia 15(1): 185-232. Keen, A. M. 1971. Sea shells of tropical west America. 1064 pp., 22 color pis. Stanford Univ. Press, Stanford, Calif. Kisch, B.-S. 1959. La collection de Chemnitzidae du Marquis de Folin au Muséum National d’Histoire Naturelle. J. Conchyliol. 99:89-112, 1 fig. Lyons, W. G. 1976. Distribution of Cerithium litteratum (Bom) (Gastropoda: Ceri­ thiidae) off western Florida. Veliger 18(4): 375-377, 1 pi. ------. 1977. Comments on three Jamaican melanellid species described by C. B. Adams (Gastropoda: Melanellidae). Occ. Papers on Mollusks, Harvard Univ. 4(55): 149-157, 1 pi. Mayr, E. 1969. Principles of systematic zoology, xi -f- 428 pp. McGraw-Hill Book Co., New York. VOLUME 91, NUMBER 2 545

Mörch, O. A. 1875. Synopsis Molluscorum marinorum Indiarum oeudentalium. Mal. Bl. 22:142-184. Thiele, J. 1929. Handbuch der Systematischen Weichtierkunde. Part 1. 376 pp. Gustav Fischer, Jena. Turgeon, D. D., and W. G. Lyons. 1978. A tropical marine molluscan assemblage in the northeastern Gulf of Mexico. (Abstract.) Bull. Am. Malacol. Union for 1977 (1978): 88-89.

Florida Department of Natural Resources, Marine Research Laboratory, 100 Eighth Avenue SE, St. Petersburg, Florida 33701.